Ranging Patterns of Two Syntopic Howler Monkey Species (Alouatta guariba and A. caraya) in Northeastern Argentina

To avoid competition, ecologically similar and closely related species tend to differ in their patterns of habitat use when they live in sympatry. We compared ranging patterns of brown howler (Alouatta guariba) and black and gold howler (A. caraya) monkeys living syntopically, i.e., co-occurring and overlapping their ranges in the same habitat within the zone of sympatry, in the Atlantic Forest of northeastern Argentina with the objective of evaluating whether their use of space contributes to the avoidance of interspecific competition for food resources. During 12 mo we collected data on the ranging behavior of 2 groups of each howler species. We analyzed annual and seasonal daily path lengths and movement rates, home range size, use and overlap, habitat and vertical strata use, and intergroup encounters. Black and gold howlers traveled farther and faster during the time of relative food abundance (abundant season) than during the time of relative food shortage (lean season), and their movement rates were affected by group identity and increased with the proportion of fruits in the diet. Brown howlers’ traveling patterns were not affected by any of these factors. Home ranges for both species (95% fixed kernel; brown howlers: 31–70 ha, black-and-gold howlers: 17–112 ha) were among the largest recorded for Alouatta. For both species, core areas (50% fixed kernel) were larger for larger versus smaller groups, and decreased in the lean season compared to the abundant season. Both species showed similar patterns of habitat use, except for a slight vertical stratification. Groups of different species overlapped their ranging areas consistently more and responded to one another less aggressively during encounters than groups of the same species, suggesting that interspecific spatial niche separation for these two syntopic species is not occurring. The vertical stratification, as well as a day-to-day avoidance strategy, may be the only responses of species to one another that could reduce the potentially high levels of competition for food suggested by their elevated trophic niche overlap. A high degree of niche overlap may explain the parapatric distribution of howlers and other closely related and ecologically similar species of primates.     

Group size in folivorous primates: ecological constraints and the possible influence of social factors

The ecological-constraints model assumes that food items occur in depletable patches and proposes that an increase in group size leads to increased day range due to more rapid patch depletion. Smaller groups become advantageous when an increase in travel costs is not repaid by an increase in energy gained or some other fitness advantage. On the other hand, we also know that group size can be influenced by social factors. Here we contrast the diet and group size of red colobus (Procolobus badius) and black-and-white colobus (Colobus guereza) in Kibale National Park, Uganda to consider how ecological and social factors are affecting their group sizes. Subsequently, we examine whether the insights gained from this detailed comparison can provide an understanding of why the social organization and group size of mantled howlers (Alouatta palliata) and black howlers (A. pigra) differ. Two groups of red colobus and two groups of black-and-white colobus were studied over 10 months. Red colobus groups were larger (48 and 24) than black-and-white colobus groups (9 and 6). The two groups of red colobus overlap home ranges with the two groups of black-and-white colobus; 75% and 95% of their home ranges were within red colobuss home range. There was a great deal of similarity in the plant parts eaten by the two species and both species fed primarily on young leaves (red colobus 70%, black-and-white colobus 76%). In terms of the actual species consumed, again there was a great deal of similarity between species. The average dietary overlap among months for the two neighboring groups of red colobus was 37.3%, while the dietary overlap between the red colobus and the black-and-white colobus group that had its home range almost entirely within the home range of the red colobus groups averaged 43.2% among months. If ecological conditions were responsible for the difference in group size between the two colobine species, one would expect the density of food trees to be lower in the home ranges of the black-and-white colobus monkeys, since they have the smaller group size. We found the opposite to be true. Both black-and-white colobus groups had more food trees and the cumulative size of those trees was greater than those in the red colobuss home ranges. We quantify how these differences parallel differences in mantled and black howlers. The average group size for mantled howlers was 12.9 individuals, and for black howlers it was 5.3 individuals. We explore possible social constraints, such as infanticide, that prevent black-and-white colobus and black howlers from living in large groups.This revised version was published online in April 2005 with corrections to the cover date of the issue.     

Roaring Behavior of Two Syntopic Howler Species (Alouatta caraya and A. guariba clamitans): Evidence Supports the Mate Defense Hypothesis

Long-distance calls are loud vocalizations involved in within and between group communication in animals. These calls may maintain cohesion with group members or communicate the ownership of valuable resources such as territory, food, or mates to individuals from other groups. In howlers (Alouatta spp.), three nonmutually exclusive hypotheses suggest that the ultimate function of roaring (howling) behavior is to protect resources from neighboring groups. The space/food defense hypothesis maintains that roars are used to gain exclusive access to food; the mate defense hypothesis holds that roars ensure exclusive access to mates; and the infanticide avoidance hypothesis states that the roaring behavior serves to avoid male takeovers and infanticide. To test these three hypotheses, we conducted a study on black-and-gold howlers (Alouatta caraya) and brown howlers (A. guariba clamitans) living in syntopy at El Pi?alito provincial park in northeast Argentina. We recorded 12 mo of data on the roaring behavior of two groups of each species, along with data on food availability and diet. Although all four groups overlapped extensively in their use of food resources during the study period, roars occurred much more frequently during interactions between groups of the same species than between groups of different species. Roaring frequency was not higher during the lean season, when high-quality food was less abundant. Howlers did not roar more frequently at home range boundaries, nor were roaring bouts spatially associated with valuable feeding resources. Males participated in all of the roaring bouts, whereas females participated in only 29% of them. Adults did not roar more frequently when there were unweaned infants in the group, as expected if roaring behavior decreases infanticide risk. These results provide no support for the food defense or infanticide avoidance hypotheses, but support predictions derived from the mate defense hypothesis.     

Relative Dominance and Resource Availability Mediate Mantled Howler (Alouatta palliata) Spatial Responses to Neighbors’ Loud Calls

Many animals generate long-distance vocalizations (“loud calls”) that have been hypothesized to mediate spatial interactions. Howls by mantled howlers (Alouatta palliata), specifically, are hypothesized to function in group spacing, but the mechanism by which this is accomplished remains unclear. In this study, I evaluated the role of naturally occurring howls in mediating mantled howler intergroup spatial interactions on Barro Colorado Island, Panama. Multivariate analyses examined relationships between social and environmental contextual variables and the likelihood that a group would approach or avoid a neighboring group’s howl. Mantled howlers were more likely to approach howling neighbors when fruit and flower availability was limited. Relative dominance was also a highly significant predictor of approach, though the relationship ran contrary to expectations. Groups were more likely to approach howling neighbors higher in the established dominance hierarchy than those lower down, possibly reflecting priority access of more dominant groups to higher quality resources. Other variables tested, including numeric odds, vocalization intensity, infanticide risk, and location of the howling group, were not significant predictors of spatial response. These findings are consistent with the assertion that howls contain identifying information and highlight the importance of including contextual variables in multivariate analyses of spatial responses to loud calls. These results also indicate potential contrasts between observational and playback studies by suggesting that howl location may play a greater role in responses to simulated intruders than to known neighbor groups, and that relative dominance may provide a more salient measure of competitive ability than numeric odds among groups with interaction histories.     

Individual participation in intergroup contests is mediated by numerical assessment strategies in black howler and tufted capuchin monkeys

Asymmetries in resource-holding potential between opposing groups frequently determine outcomes of intergroup contests. Since both numerical superiority and high intergroup dominance rank may confer competitive advantages, group members should benefit from assessing the relative strength of rivals prior to engaging in defensive displays. However, differences in individual assessment may emerge when cost–benefit trade-offs differ among group members. We examine the influence of numerical superiority and intergroup dominance relationships on individual participation in intergroup encounters in black howler monkeys (Alouatta pigra) and tufted capuchin monkeys (Sapajus nigritus). Black howlers responded with longer vocal displays during encounters with neighbours with an equal number of resident males, while tufted capuchins increased their participation with increasing relative male group size. Within each species, males and females responded similarly to varying numerical odds, suggesting that despite pay-off asymmetries between males and females, both sexes were similarly influenced by numerical asymmetries in deciding to participate in collective group defence. Whereas the outcome of contests among tufted capuchins was determined by relative male group size, reflected in a pronounced intergroup dominance hierarchy, the absence of dominance relationships among black howler groups may have provoked prolonged vocal displays in order to assess rival groups with matching competitive abilities.     

Social Relationships Among Male <Emphasis Type="Italic">Alouatta pigra</Emphasis>

We investigated male social relationships in 2 groups of black howlers (Alouatta pigra) in Palenque National Park, Mexico, over a 14-mo study characterized by frequent changes in male group membership. Both single males and pairs of males entered our focal groups. Single males tended to join groups, whereas pairs of males entering groups together successfully evicted resident males. The 19 male dyads across the 11 periods defined by changes in group membership were rarely in close proximity or interacting with one another. Nevertheless, 1 male formed significantly closer associations with resident females in 6 periods and achieved higher mating success during 4 of the periods when ≥1 female was sexually active. In the other 5 periods, no particular male maintained significantly closer associations with resident females, which may be a result of the rapid sequence of changes in male membership and corresponding social instability. Resident males participated frequently and consistently in mutual howling, but the resident male with the strongest female associations and highest mating success initiated howling bouts more frequently in all but 1 of the 6 socially stable periods. Though still preliminary, our findings suggest that variation in social relationships among male black howlers may be more related to their ability to establish strong relationships with females than to their relationships with one another.     

Group encounters and territoriality in wild Alaotran gentle lemurs (Hapalemur griseus alaotrensis)

During a 3 month field study, 18 group encounters between four groups of Alaotran gentle lemurs (Hapalemur griseus alaotrensis) were observed in the Lake Alaotra marshland in Madagascar. Behaviors observed during group encounters are described, and quantitative data on intergroup interactions and ranging patterns are discussed in the context of territoriality. Intergroup interactions varied in their intensity, ranging from visual monitoring, scent marking, and display locomotion to penetrating the neighboring range. A quarter (27%) of the encounters were aggressive, involving chases and confrontation displays, and no affiliative interactions were observed. Both adult males and females were involved in intergroup encounters, with males playing more active roles. All encounters took place in the small overlapping areas of neighboring home ranges. The outcome of an aggressive encounter was determined by its location, with the resident group always driving out the intruding one. The Alaotran gentle lemur actively advertised and consistently defended a large area of its home range against intrusion of neighboring groups and can therefore be considered territorial. Am. J. Primatol. 46:251–258, 1998. © 1998 Wiley-Liss, Inc.     

Aspects of Male Competition in <Emphasis Type="Italic">Colobus vellerosus</Emphasis>: Preliminary Data on Male and Female Loud Calling,and Infant Deaths After a Takeover

Male Colobus vellerosus are the main participants in intergroup encounters, and lead incursions in neighboring groups during which they attack infants. Extragroup copulations, all-male groups, and male takeover occur in the species. Here, we provide additional information on behaviors associated with male reproductive competition in Colobus vellerosus. We examined 1 resident male loud calling and participation in intergroup encounters in relation to a takeover. We also report a second case of takeover that led to the death of the former resident male and the death of 2 male infants, presumably as a result of aggression from the all-male group. The new resident male wounded the third infant of the group, which apparently died after its mother abandoned it. During the period characterized by the attacks on the infant and after its disappearance, females initiated and participated in loud call bouts with the new resident male. We examine the possible functions of female loud calling, and suggest that in this context, it might force the resident male to call along to indicate his presence.     

Experimental test of female black howler monkey (Alouatta Pigra) responses to loud calls from potentially infanticidal males: effects of numeric odds, vulnerable offspring, and companion behavior

During group defense, the contribution of female black howler monkeys (Alouatta pigra) may help deter male intruders; however, their involvement during natural intergroup encounters is facultative. Experimental playback trials simulating potentially infanticidal males were used to examine whether a female's reproductive investment and/or her group's relative fighting ability would influence her participation in loud call displays. Female howlers never responded to recordings without alpha male accompaniment, but their response patterns did not simply mimic his. For example, unlike alpha males, females with small, vulnerable offspring were no more likely than females without infants to participate in howling displays during playback trials. Further, although females without any small infants in their group got closer to speakers than females living in groups with small infants, males did not respond in the same way. To artificially simulate different "numeric odds" scenarios, recordings of one or three howling males (simulated intruders) were broadcast to females living in groups with 1-3 resident males. Responses were consistent with the hypothesis that females assessed intergroup fighting ability. As in alpha males, the weakest female responses occurred when the numeric odds were against their group. However, whereas alpha males participated most when numeric odds were in their favor, females had the most intense approach responses when the number of defending and intruding males was equal. Females appeared to use a cost-effective strategy, reserving their assistance for when their participation could have the greatest impact.     

The Function of Howling in the Ring-Tailed Lemur (Lemur catta)

Long calls are sex-specific vocalizations used for mate attraction or mate defense in many animal species. Ring-tailed lemurs (Lemur catta), female-dominant strepsirrhines, have a male-specific long call termed a howl, with proposed functions that have never been empirically tested. I aimed to investigate why ring-tailed lemur males howl and to test whether the mate defense and mate attraction hypotheses for long-calling were applicable to this species. From March to July 2010, I collected 600 h of focal data on 25 males aged ≥3 year at Beza Mahafaly Special Reserve, Madagascar. I observed each male continuously for 30 min at a time and noted all agonism using one–zero sampling at 2.5-min intervals. I calculated male dominance rank from these data. I recorded days when female estrus occurred and noted howling and intergroup encounters using all-occurrences sampling. Howling rate was not significantly related to female estrus or male dominance rank, providing no support for the mate attraction hypothesis or the intragroup mate defense hypothesis. In contrast, the intergroup mate defense hypothesis was strongly supported. During intergroup encounters, male howling rate significantly increased compared to howling rate at times without other groups present, and a greater number of males participated in multimale howling choruses when compared to times without nongroup members present. My results suggest that male ring-tailed lemurs howl to advertise their presence and location to other groups, but not to male or female members of their own group. Howling could discourage male immigration by advertising the number of males already present in a group. Long calls are used for similar mate defense purposes during intergroup encounters by other primates, including Thomas langurs (Presbytis thomasi) and chacma baboons (Papio ursinus).     

Intergroup relations and home range use in Verreaux's sifaka (Propithecus verreauxi)

Relationships between neighboring groups feature prominently in socioecological theory, but few empirical studies have focused on the effects of neighbors on the behavior of primates. Interactions between neighboring groups are most conspicuous during intergroup encounters, but the likelihood of encounters with neighbors can also affect ranging and activity patterns indirectly, and, as a result, behavioral patterns in areas of exclusive use may differ from those in overlap areas of adjacent home ranges. The aim of this study was to examine intergroup relations and spatial variation in home range use in Verreaux's sifaka (Propithecus verreauxi) during the annual mating season. Based on 230 hr of focal animal data collected from ten members of five adjacent groups, we found that behavioral patterns and patterns of resource utilization were not markedly different between areas of exclusive use and overlap areas of adjacent home ranges. Group cohesion tended to be higher in overlap than in core areas, but the proportion of time spent resting and foraging did not differ between these two areas. However, dominant males exhibited a higher scent-marking rate in overlap areas. Observations during intergroup encounters revealed that chases between males occurred frequently, whereas fights involving physical contact were not observed. We also found that the intergroup dominance hierarchy depended on the relative group size or the number of males in each group, with groups of lower dominance rank exhibiting a tendency to sleep proportionally more often in their core areas. The results of this study suggest that in primate species with a moderate level of intergroup aggression, such as Verreaux's sifaka, the possibility of encountering neighbors in areas of home range overlap does not exert strong influence on their behavior and resource utilization.     

Who is coordinating collective movements in black and gold howler monkeys?

Decisions about when and where to travel are likely to have a strong influence on the feeding, ecology, and foraging strategies of individual primates living in a cohesive social group. Specifically, given differences in age, sex, reproductive status, or social dominance, particular group members may benefit from remaining at their present location while others may benefit from traveling to another area of their range to feed or rest. In this study, we present data on movement coordination in two groups of wild black and gold howler monkeys inhabiting Isla Brasilera (27º 20′S and 58º 40′W) in northern Argentina. We examine how factors such as sex, age, reproductive status, and dominance affect patterns of group movement coordination at feeding or resting sites, and in the context of intergroup encounters. Two groups were followed five days a month from sunrise to sunset during June to November 2004. Using focal and scan sampling techniques, we recorded 262 group displacements, the identity of the individual initiating and leading displacement, and the identity of the first individual to arrive at feeding, resting, or intergroup encounter sites. We found that overall age was the only factor that influenced group coordination: adults led more often (94.5 %) than immature individuals (5.5 %) in both groups. We did not find differences among adults. However, we found that males lead more often than females at intergroup encounters, consistent with the male-mate defense hypothesis. The distributed leadership pattern among adults observed in this study may suggest that adult individuals make equally shared consensus decisions. This pattern should be further examined using this individual-level approach in other populations of black and gold howlers, other species of howlers, and in other atelines in which within-group social tolerance is the rule rather than the exception.     

Role of loud calls in brown howlers,Alouatta fusca

The loud calls of brown howler monkeys were studied during a year at the Santa Genebra Reserve, in southeastern Brazil. The study group emitted roars and barks on a total of 47 occasions, the majority of which (92%) were restricted to intergroup visual encounters. Loud calls were also elicited by the roars of distant groups (6%) and during intragroup agonistic interactions (2%). Intergroup visual encounters (n = 42) occurred predominantly in seldom used quadrats of the study group home range. In these instances, the loud calls were produced chiefly by the adult male alone (69% of cases), while the study group's two adult females joined the male in the remaining cases. Intergroup physical aggression, such as chase and displacement, was observed during 15 encounters (35% of cases). A dawn chorus does not occur in Santa Genebra—the loud calls were heard most frequently in mid-morning and again during mid-afternoon—and they were more abundant during the dry season, when the availability of food (new leaves) in the forest was lower. The data presented here provide some support for the hypothesis that roars of howler adult males are used in assessment of opponents, providing an alternative to energetically expensive chases and fights. However, given the relatively high rate of physical aggression observed during intergroup encounters, a result probably related to the high density of howlers and the consequent high frequency of intergroup encounters observed in this forest (0.7/day), ritualized aggression, in the form of loud calling, is apparently often insufficient to settle disputes. © 1995 Wiley-Liss, Inc.     

When Top-Down Becomes Bottom Up: Behaviour of Hyperdense Howler Monkeys (Alouatta seniculus) Trapped on a 0.6 Ha Island

Predators are a ubiquitous presence in most natural environments. Opportunities to contrast the behaviour of a species in the presence and absence of predators are thus rare. Here we report on the behaviour of howler monkey groups living under radically different conditions on two land-bridge islands in Lago Guri, Venezuela. One group of 6 adults inhabited a 190-ha island (Danto) where they were exposed to multiple potential predators. This group, the control, occupied a home range of 23 ha and contested access to food resources with neighbouring groups in typical fashion. The second group, containing 6 adults, was isolated on a remote, predator-free 0.6 ha islet (Iguana) offering limited food resources. Howlers living on the large island moved, fed and rested in a coherent group, frequently engaged in affiliative activities, rarely displayed agonistic behaviour and maintained intergroup spacing through howling. In contrast, the howlers on Iguana showed repulsion, as individuals spent most of their time spaced widely around the perimeter of the island. Iguana howlers rarely engaged in affiliative behaviour, often chased or fought with one another and were not observed to howl. These behaviors are interpreted as adjustments to the unrelenting deprivation associated with bottom-up limitation in a predator-free environment.     

Can a group elicit duets from its neighbours? A field study on the black-crested gibbon (Nomascus concolor jingdongensis) in Central Yunnan, China

We tested the intergroup spacing hypothesis with a 13-month field study of the interaction of singing behaviour between 3 neighbouring groups of black-crested gibbons (Nomascus concolor jingdongensis) at Mt. Wuliang, Central Yunnan, China. Neighbouring groups tended not to sing on the same day. While it did happen occasionally, a group seldom started a duet while its neighbour was singing, or within 5 min of the singing ceasing. The intersong intervals of 2 groups calling on the same day were similar or significantly longer than the intervals between any song bouts randomly selected from the same 2 groups when calling on different days. Groups did not sing in intergroup encounters and showed similar responses to neighbouring groups and groups with no common border. These results did not support the intergroup spacing hypothesis. The possible reasons are discussed.     

Dispersal patterns in Costa Rican mantled howling monkeys

Both male and female juveniles disperse in Costa Rican mantled howling monkeys (Alouatta palliata). 79% of the males and 96% of the females leave their natal groups. Males may spend up to 4 years and females up to 1 year as solitaries. Extra-group individuals are faced with only three possibilities, i.e., form a new group by joining another extra-group individual, join an established social group, or remain solitary. Most surviving extra-group individuals join an established social group which contains no kin. Females join with the help of a resident male and once in a group proceed to rise to the alpha position through dyadic interactions. The immigrant female either becomes the alpha female or leaves and tries again in another group. Males challenge the alpha male and either defeat him or remain solitary. Competition with relatives for limited high quality food may be the reason for both sexes leaving their natal groups in howlers. By leaving, the successful immigrants increase their mothers inclusive fitness while suppressing the fitness of nonrelatives instead of remaining natal and competing with relatives for limited food.     

Against all odds: Numerical assessment by tufted capuchin monkeys

When competitors are able to assess the asymmetry in the resource holding potential before interacting, individuals or groups should avoid interacting with stronger opponents, thereby avoiding the energy costs and risk of injury associated with aggressive intergroup encounters. Thus, escalated aggression is expected only between closely matched competitors. Among Argentine tufted capuchin monkeys (Sapajus nigritus), intergroup dominance is decided by the asymmetry in male group size. Using playback experiments, I simulated intergroup encounters with neighboring groups, manipulating both the apparent numerical asymmetry and the resource context. During experimental trials, I recorded the approach behavior of the focal individual, as well as changes in neighbor density and individual travel speed following the presentation of the playback stimulus, to assess whether individual willingness to participate in resource defense was affected by the probability of winning the encounter. In spite of the competitive disadvantage, neither males nor females showed a decreased probability of approach when the numerical odds strongly favored the opposing group. Instead decisions regarding whether to participate appear to be driven primarily by the resource context. Nevertheless, changes in individual behavior during approaches suggest that tufted capuchin monkeys are sensitive to the relative odds. Individuals accelerated less when approaching a larger group, although no changes in neighbor density were apparent. The absence of an effect of the numerical asymmetry on willingness to approach the playback speaker suggests that subordinate groups benefit from engaging in intergroup aggression with larger neighbors, despite the high probability of losing. These encounters may serve to assess the current subjective resource valuation of the neighboring group or limit territorial expansion by large groups by decreasing the marginal value of home range exclusivity. Because these encounters are riskier, however, individuals appear to alter their approaches, becoming more tentative as the numerical odds increasingly favor the opposing group.     

Causes of Intraspecific Aggression in Alouatta palliata mexicana: Evidence from Injuries,Demography, and Habitat

Although howlers have been traditionally considered to be pacific, showing one of the lowest rates of aggression among primates, new evidence is emerging to question this image. We present data on injuries in Mexican mantled howlers (Alouatta palliata mexicana) in relation to different sociecological parameters. We censused howler populations in 19 forest fragments over a 17-mo period in the Los Tuxtlas Region, southern Veracruz, Mexico. We conducted detailed observations of scars, torn lips, broken fingers, mutilated tails and other visible injuries. We also collected data on the demography, biogeography and vegetation of each fragment in order to relate injury data to them. We censused 333 howlers, of which we exhaustively observed 254 for injuries. Four resident adult females (n = 108) and 29 adult resident males (n = 76) had injuries, while none of the solitary males (n=16), solitary females (n=1), juveniles (n=23) and infants (n=30) had them. We discuss possible interpretative scenarios for the distribution of injuries. Although some results suggest that food resource concentration may determine intergroup agonistic encounters, we propose that physical injury is primarily associated with male-male agonistic encounters during takeovers, and consequently it could indicate migration among troops.     

Diet, foraging, and use of space in wild golden-headed lion tamarins

Lion tamarins (Callitrichidae: Leontopithecus) are small frugi-faunivores that defend large home ranges. We describe results from the first long-term investigation of wild golden-headed lion tamarins (L. chrysomelas; GHLTs). We present data about activity budgets, daily activity cycles, diet, daily path length, home range size, home range overlap, and territorial encounters for three groups of GHLTs that were studied for 1.5-2.5 years in Una Biological Reserve, Bahia State, Brazil, an area characterized by aseasonal rainfall. We compare our results to those from other studies of lion tamarins to identify factors that may influence foraging and ranging patterns in this genus. Ripe fruit, nectar, insects, and small vertebrates were the primary components of the GHLT diet, and gums were rarely eaten. Fruit comprised the majority of plant feeding bouts, and the GHLTs ate at least 79 different species of plants from 32 families. The most common foraging sites for animal prey were epiphytic bromeliads. The GHLTs defended large home ranges averaging 123 ha, but showed strong affinities for core areas, spending 50% of their time in approximately 11% of their home range. Encounters with neighboring groups averaged two encounters every 9 days, and they were always aggressive. Data about time budgets and daily activity cycles reveal that the GHLTs spent most of their time foraging for resources or traveling between foraging sites distributed throughout their home ranges. The GHLTs spent much less time consuming exudates compared to lion tamarins in more seasonal environments. Additionally, the GHLTs had much larger home ranges than golden lion tamarins (L. rosalia), and did not engage in territorial encounters as frequently as L. rosalia. GHLT ranging patterns appear to be strongly influenced by resource acquisition and, to a lesser extent, by resource defense.     

Howling activity of free-ranging wolves (<Emphasis Type="Italic">Canis lupus</Emphasis>) in the Białowieża Primeval Forest and the Western Beskidy Mountains (Poland)

We investigated spontaneous howling by radio-collared wolves Canis lupus inhabiting the Białowieża Primeval Forest (BPF), eastern Poland, and elicited howling behavior in wolves of the BPF and the Western Beskidy Mountains, southern Poland. Over half (58%) of all spontaneous howls recorded throughout a year occurred in the period from July to October, with a peak in August. The daily pattern of vocal activity by wolves was characterised by a peak between 1800 and 0000 hours, which coincided with the first (dusk) peak of wolf mobility. Wolves howled from the core areas of their territories and not from the peripheries. Howls served as communication between temporarily separated pack mates (43% of cases), after re-union (18%) and before setting out for a hunt (22%). Very few spontaneous howls (2%) were targeted at a neighbouring pack. Wolves responded to human-simulated howling in June–September, with a peak in August (reply rate: 39%). The duration of elicited howling increased significantly with group size: howls by single wolves or pairs lasted, on average, 34–40 s, whereas those of five to seven wolves (including pups) had an average duration of 67–95 s, with a maximum length of nearly 4 min. In the populations of Polish wolves studied here, spontaneous howling served primarily for intra-pack communication. Nonetheless, the high reply rate to howling simulation showed that – if necessary – wolves readily advertised their presence in a territory to strangers.